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PubMed2002

Audiogram of a harbor porpoise (Phocoena phocoena) measured with narrow-band frequency-modulated signals.

The underwater hearing sensitivity of a two-year-old harbor porpoise was measured in a pool using standard psycho-acoustic techniques. The go/no-go response paradigm and up-down staircase psychometric method were used. Auditory sensitivity was measured by using narrow-band frequency-modulated signals having center frequencies between 250 Hz and 180 kHz. The resulting audiogram was U-shaped with the range of best hearing (defined as 10 dB within maximum sensitivity) from 16 to 140 kHz, with a reduced sensitivity around 64 kHz. Maximum sensitivity (about 33 dB re 1 microPa) occurred between 100 and 140 kHz. This maximum sensitivity range corresponds with the peak frequency of echolocation pulses produced by harbor porpoises (120-130 kHz). Sensitivity falls about 10 dB per octave below 16 kHz and falls off sharply above 140 kHz (260 dB per octave). Compared to a previous audiogram of this species (Andersen, 1970), the present audiogram shows less sensitive hearing between 2 and 8 kHz and more sensitive hearing between 16 and 180 kHz. This harbor porpoise has the highest upper-frequency limit of all odontocetes investigated. The time it took for the porpoise to move its head 22 cm after the signal onset (movement time) was also measured. It increased from about 1 s at 10 dB above threshold, to about 1.5 s at threshold.

Animals↗
PubMed2003

The mammalian cochlear map is optimally warped.

The form of the mammalian cochlear frequency-position map has been well described by Greenwood and empirical values found for its coefficients for a number of species. The apical portion of the mammalian map is spatially compressed relative to the base, and this nonuniformity in the representation of frequency is evidently consistent across species. However, an evolutionary reason for this consistency, encompassing critical band behavior with respect to position, is conspicuously missing. Likewise, the length of the cochlea in any mammal, including echolocating species, is related to body size, but attempts to explain the length in terms of frequency limits, range, or resolution have no general explanation. New insight stems from a hypothesis in which the map curvature may be appreciated as an adaptation for optimal frequency resolution over the auditory range. It is demonstrated numerically that the mammalian curve may be considered a member of a family of curves which vary in their degree of warp. The "warp factor" found to be common across mammals is an optimal trade-off between four conflicting constraints: (1) enhancing high-frequency resolution; (2) setting a lower bound on loss of existing low-frequency resolution; (3) minimizing map nonuniformity; and (4) keeping the whole map smooth, thereby avoiding reflections.

Animals↗
PubMed2003

The sonar beam pattern of a flying bat as it tracks tethered insects.

This paper describes measurements of the sonar beam pattern of flying echolocating bats, Eptesicus fuscus, performing various insect capture tasks in a large laboratory flight room. The beam pattern is deduced using the signal intensity across a linear array of microphones. The positions of the bat and insect prey are obtained by stereoscopic reconstruction from two camera views. Results are reported in the form of beam-pattern plots and estimated direction of the beam axis. The bat centers its beam axis on the selected target with a standard deviation (sigma) of 3 degrees. The experimental error is +/- 1.4 degrees. Trials conducted with two targets show that the bat consistently tracks one of the targets with its beam. These findings suggest that the axis of the bat sonar beam is a good index of selective tracking of targets, and in this respect is analogous to gaze in predominantly visual animals.

Animals↗
PubMed2004

The bat head-related transfer function reveals binaural cues for sound localization in azimuth and elevation.

Directional properties of the sound transformation at the ear of four intact echolocating bats, Eptesicus fuscus, were investigated via measurements of the head-related transfer function (HRTF). Contributions of external ear structures to directional features of the transfer functions were examined by remeasuring the HRTF in the absence of the pinna and tragus. The investigation mainly focused on the interactions between the spatial and the spectral features in the bat HRTF. The pinna provides gain and shapes these features over a large frequency band (20-90 kHz), and the tragus contributes gain and directionality at the high frequencies (60 to 90 kHz). Analysis of the spatial and spectral characteristics of the bat HRTF reveals that both interaural level differences (ILD) and monaural spectral features are subject to changes in sound source azimuth and elevation. Consequently, localization cues for horizontal and vertical components of the sound source location interact. Availability of multiple cues about sound source azimuth and elevation should enhance information to support reliable sound localization. These findings stress the importance of the acoustic information received at the two ears for sound localization of sonar target position in both azimuth and elevation.

Animals↗
PubMed2006

Auditory brainstem response in a harbor porpoise show lack of automatic gain control for simulated echoes.

The auditory brainstem response (ABR) response to simulated echolocation clicks was studied in a harbor porpoise, Phocoena phocoena, to determine the relationship between the animal's perceived echo strength and the simulated target distance. In one experiment the click level at the listening post was kept constant while delay was changed, in another, the level was varied to approximate spreading losses. Results of both experiments indicate that there is no automatic gain control in the hearing system of this harbor porpoise.

Animals↗
PubMed2006

Comment on paper entitled, "An inversion of Freedman's 'image pulse' model in air". Acoust. Soc. Am. 119(2), 965-975 (2006).

Echolocation (i.e., perceiving objects using acoustic echoes) is well-known in underwater detection and to a lesser extent in robot guidance and machine perception. The paper by Tsakiris and McKerrow is concerned with machine perception in air using Freedman's asymptotic model, which was originally developed to predict the backscattering multiple-echo effect observed in sonar detection. This effect was subsequently shown to be due to the elastic response of underwater targets. Freedman's model can be used in air because the acoustic target is assumed to be rigid. Also, the model's prediction of multiple echoes can be used to obtain information about the shape of the target. This is the so-called inversion of the Freedman model by Tsakiris and McKerrow. In their paper, various simple bodies are tested in air using ultrasound and it is shown that the model provides relatively poor information about body shape. Several explanations are given. However, one explanation is not considered, namely that the model itself is not satisfactory. First, there is poor agreement with exact backscattering theory. Second, deriving information about target shape from the multiple echoes predicted by the model is a highly questionable procedure. Both these aspects are examined here.

Acoustics↗
PubMed1986

Accuracy of distance measurement in the bat Eptesicus fuscus: theoretical aspects and computer simulations.

Behavioral experiments of Simmons [J. Acoust. Soc. Am. 54, 157-173 (1973) and Science 204, 1336-1338 (1979)] on the ranging accuracy in the bat Eptesicus fuscus have led to far-reaching postulates on the existence of optimal and phase-conserving processing mechanisms in the bat. In this paper, the results of computer simulations of these experiments are presented. Two receiver types are investigated: the fully coherent cross-correlation receiver and the cross-correlation receiver with envelope processing (semicoherent). It is shown that Simmons' experiments cannot be treated as a simple estimation of distance, but require at least two (range difference experiment; see Simmons, 1973) or four (range jitter experiment; see Simmons, 1979) echolocation sounds for one decision. The performance of the bat in both experiments is much worse than predicted for a coherent and a semicoherent receiver type. The bat's accuracy in Simmons' range difference experiment is at least 18 dB worse than predicted for an optimal receiver. The results of the jitter experiment cannot be interpreted in a simple way as proof that bats are able to evaluate phase information as in a fully coherent cross-correlation receiver.

Animals↗
PubMed1986

Target detection by the beluga using a surface-reflected path.

During an echolocation-in-noise experiment we suspected that a beluga (Delphinapterus leucas) was using a surface-reflected path to maximize detection performance. We tested and confirmed this suspicion by acoustically and mechanically denying access to surface path information. The whale's performance varied as the surface-reflected path was denied.

Animals↗
PubMed1989

Insights into dolphin sonar discrimination capabilities from human listening experiments.

A variety of dolphin sonar discrimination experiments have been conducted, yet little is known about the cues utilized by dolphins in making fine target discriminations. In order to gain insights on cues available to echolocating dolphins, sonar discrimination experiments were conducted with human subjects using the same targets employed in dolphin experiments. When digital recordings of echoes from targets ensonified with a dolphinlike signal were played back at a slower rate to human subjects, they could also make fine target discriminations under controlled laboratory conditions about as well as dolphins under less controlled conditions. Subjects reported that time-separation-pitch and duration cues were important. They also reported that low-amplitude echo components 32 dB below the maximum echo component were usable. The signal-to-noise ratio had to be greater than 10 dB above the detection threshold for simple discrimination and 30 dB for difficult discrimination. Except for two cases in which spectral cues in the form of "click pitch" were important, subjects indicated that time-domain rather than frequency-domain processing seemed to be more relevant in analyzing the echoes.

Animals↗
PubMed1994

The brain-stem auditory-evoked response in the big brown bat (Eptesicus fuscus) to clicks and frequency-modulated sweeps.

Three experiments were performed to evaluate the effects of stimulus level on the brain-stem auditory-evoked response (BAER) in the big brown bat (Eptesicus fuscus), a species that uses frequency-modulated (FM) sonar sounds for echolocation. In experiment 1, the effects of click level on the BAER were investigated. Clicks were presented at levels of 30 to 90 dB pSPL in 10-dB steps. Each animal responded reliably to clicks at levels of 50 dB pSPL and above, showing a BAER containing four peaks in the first 3-4 ms from click onset (waves i-iv). With increasing click level, BAER peak amplitude increased and peak latency decreased. A decrease in the i-iv interval also occurred with increasing click level. In experiment 2, stimuli were 1-ms linear FM sweeps, decreasing in frequency from 100 to 20 kHz. Stimulus levels ranged from 20 to 90 dB pSPL. BAERs to FM sweeps were observed in all animals for levels of 40 dB pSPL and above. These responses were similar to the click-evoked BAER in waveform morphology, with the notable exception of an additional peak observed at the higher levels of FM sweeps. This peak (wave ia) occurred prior to the first wave seen at lower levels (wave ib). As the level of the FM sweep increased, there was a decrease in peak latency and an increase in peak amplitude. Similarity in the magnitude and behavior of the i-iv and ib-iv intervals suggests that wave ib to FM sweeps is the homolog of the wave i response to click stimuli. Experiment 3 tested the hypothesis that wave ia represented activity emanating from more basal cochlear regions than wave ib. FM sweeps (100-20 kHz) were presented at 90 dB pSPL, and broadband noise was raised in level until the BAER was eliminated. This "masked threshold" occurred at 85 dB SPL of noise. At masked threshold, the broadband noise was steeply high-pass filtered at five cutoff frequencies ranging from 20 to 80 kHz. Generally, wave ia was eliminated for masker cutoff frequencies of 56.6 kHz and below, while wave ib was typically observed for masker cutoffs down to 28.3 kHz. The results of these three experiments are compared and contrasted with data from other mammalian BAER studies.

Acoustic Stimulation↗
PubMed1995

Spatially dependent acoustic cues generated by the external ear of the big brown bat, Eptesicus fuscus.

To measure the directionality of the external ear of the echolocating bat, Eptesicus fuscus, the left or right eardrum of a dead bat was replaced by a microphone which recorded signals received from a sound source that was moved around the stationary head. The test signal was a 0.5-ms FM sweep from 100 kHz to 10 kHz (covering all frequencies in the bat's biosonar sounds). Notches and peaks in transfer functions for 7 tested ears varied systematically with changes in elevation. For the most prominent notch, center frequency decreased from about 50 kHz for elevations at or near the horizontal to 30-40 kHz for elevations 30 degrees-40 degrees below the horizontal. A second notch shifted from about 85 kHz to 70 kHz over these same elevations. Above the horizontal, a peak that flanks these notches changed in amplitude by 15 dB with changes in elevation. Removal of the tragus from the external ear disrupted the systematic movement of notch frequencies with elevation but did not disrupt changes in the peak's amplitude. Smaller changes in notch frequency also occurred with changes in azimuth, so monaural notch information alone cannot determine the position of sound sources away from the median plane. However, because bats routinely keep the head pointed at the target's azimuth, median-plane localization occurs with monaural cues delivered to the two ears. Corresponding changes with elevation occurred in the impulse-response, which consists of a series of 3-6 peaks spaced 10-20 microseconds apart. The time separation of two prominent impulse peaks systematically increased from 22-26 microseconds above the horizontal to about 36-40 microseconds below the horizontal, and removal of the tragus disrupted this time shift below the horizontal.

Animals↗
PubMed1995

Neural network modeling of a dolphin's sonar discrimination capabilities.

The capability of an echolocating dolphin to discriminate differences in the wall thickness of cylinders was previously modeled by a counterpropagation neural network using only spectral information from the echoes. In this study, both time and frequency information were used to model the dolphin discrimination capabilities. Echoes from the same cylinders were digitized using a broadband simulated dolphin sonar signal with the transducer mounted on the dolphin's pen. The echoes were filtered by a bank of continuous constant-Q digital filters and the energy from each filter was computed in time increments of 1/bandwidth. Echo features of the standard and each comparison target were analyzed in pairs by a counterpropagation neural network, a backpropagation neural network, and a model using Euclidean distance measures. The backpropagation network performed better than both the counterpropagation network, and the Euclidean model, using either spectral-only features or combined temporal and spectral features. All models performed better using features containing both temporal and spectral information. The backpropagation network was able to perform better than the dolphins for noise-free echoes with Q values as low as 2 and 3. For a Q of 2, only temporal information was available. However, with noisy data, the network required a Q of 8 in order to perform as well as the dolphin.

Animals↗
PubMed1996

Sound source elevation and external ear cues influence the discrimination of spectral notches by the big brown bat, Eptesicus fuscus.

Measurements of external ear transfer functions in the echolocating bat Eptesicus fuscus have revealed a prominent spectral notch that decreases in center frequency (50 to 30-35 kHz) as elevation decreases [Wotton et al., J. Acoust. Soc. Am. 98, 1423-1445 (1995)]. To examine the influence of this notch, four Eptesicus were trained to discriminate between two sets of electronically generated artificial echoes. The negative (unrewarded) stimulus contained a test spectral notch at a specific frequency that varied from 30 to 50 kHz, while the positive (rewarded) stimulus contained no test notch. The vertical position of the loudspeakers delivering these simulated echoes was changed daily. When echoes were returned from an elevation at which the external ear introduced a spectral notch at the same frequency as the test notch, then the discrimination should have been difficult. The bats' performance conformed to this prediction: All bats discriminated the presence of a 35-kHz notch at all elevations except -10 degrees. As the frequency of the synthesized notch increased, the elevation at which bats could not perform the discrimination also increased. The movement of the bat's "blind spot" for the test notch of different frequencies followed the movement of the external ear notch at different elevations.

Animals↗
PubMed1997

Hearing deficits measured in some Tursiops truncatus, and discovery of a deaf/mute dolphin.

Eight bottlenose dolphins Tursiops truncatus (four male, four female) were trained to respond to 100-ms tones. Three male dolphins (ages 23, 26, and 34) exhibited hearing disability at four higher frequencies-70, 80, 100, and 120 kHz even at 111-135 dB re: 1 microPa. Two females (ages 32 and 35) responded to all frequencies as did a male (age 7) and a female (age 11). One female (age 33) responded to all tones at 80 kHz and below; however, she failed to respond at 100 or 120 kHz. One young female dolphin (age 9) exhibited no perception of sound to behavioral or electrophysiological tests. This young female was not only deaf, but mute. The dolphin was monitored periodically by hydrophone and daily by trainers (by ear in air) for 7 years until she was age 16. The animal never whistled or made echolocation pulses or made burst pulse sounds as other dolphins do.

Age Factors↗
PubMed1998

Underwater audiogram of a tucuxi (Sotalia fluviatilis guianensis).

Using a go/no go response paradigm, a tucuxi (Sotalia fluviatilis guianensis) was trained to respond to pure-tone signals for an underwater hearing test. Auditory thresholds were obtained from 4 to 135 kHz. The audiogram curve shows that this Sotalia had an upper limit of hearing at 135 kHz; from 125 to 135 kHz sensitivity decreased by 475 dB/oct. This coincides with results from electrophysiological threshold measurements. The range of best hearing (defined as 10 dB from maximum sensitivity) was between 64 and 105 kHz. This range appears to be narrower and more restricted to higher frequencies in Sotalia fluviatilis guianensis than in other odontocete species that had been tested before. Peak frequencies of echolocation pulses reported from free-ranging Sotalia correspond with the range of most sensitive hearing of this test subject.

Animals↗
PubMed1998

Signal processing of the echo signatures returned by submerged shells insonified by dolphin "clicks:" active classification.

A large set of dolphin-emitted acoustic pulses ("echolocation clicks") have been examined, which were reflected from various elastic shells that were suspended, underwater, 4.5 m in front of the animal in a large test site in Kaneohe Bay, Hawaii. A carefully instrumented analog-to-digital system continuously captured the emitted clicks and also the returned, backscattered echoes (A/D conversion at 500 kHz). Using standard conditioning techniques and food reinforces, the dolphin is taught to push an underwater paddle when the "correct" target-the one he has been trained to identify-is presented to him. He communicates his consistently correct identifying choices in this manner. Many echoes returned by three types of cylindrical shells in both the time and frequency domains as well as in the joint time-frequency (t-f) domain, by means of Wigner-type distributions have been examined. It will be shown exactly how specific features observable in these displays are directly related to the physical characteristics of the shells. This processing takes advantage of certain fundamental resonance principles to show which echo features contain information about the size, shape, wall thickness, and material composition of both the shell and its filler substance. In the same fashion that these resonance features give the identifying characteristics of each shell, it is believed they may also give them to the dolphin. These echo features may allow him to extract the target properties by inspection without any need for computations. It is claimed that this may be the fundamental physical explanation of the dolphin's amazing target ID feats, upon which they base their recognition choices. This claim may be substantiated by the detailed analysis of many typical echoes returned by various shells, when they are interrogated by several dolphins. Thus far, this analysis of many echoes from many shells has only been carried out for a single dolphin.

Animals↗
PubMed1998

Spontaneous classification of complex tones at high and ultrasonic frequencies in the bat, Megaderma lyra.

Megaderma lyra, a bat species using harmonically structured calls for echolocation, exploits the spectral content of its echoes for texture discrimination. It is the aim of the present study to test according to which sensory qualities harmonic complex tones are spontaneously classified by this bat. The applied experimental paradigm is especially adapted to the preference of M. lyra to use absolute pitch cues. Three animals were trained in a 2-AFC procedure to classify three-component stimuli as low or high, with all their harmonics below or above a pure tone reference of 33 kHz, respectively. Later, the original tones were interspersed with "incomplete" test stimuli, with their fundamentals (and lower harmonics) missing. These were ambiguous in that their possible virtual, i.e., collective pitches were below the reference whereas their pure tone pitches were above it. Bat 1 classified 22 of 23 test stimuli with missing fundamentals between 5.3 and 28.3 kHz according to their collective pitches, whereas bat 2 judged all presented ambiguous tones on the basis of their pure tone pitches. Bat 3 failed the pitch control criterion which is why in this case results cannot be interpreted unequivocally. The implications of these findings are discussed with respect to the bats' behavioral context, as well as to psychoacoustical models of the formation of the pitch of complex tones.

Animals↗
PubMed2000

Range discrimination by big brown bats (Eptesicus fuscus) using altered model echoes: implications for signal processing.

The sonar emissions of two big brown bats (Eptesicus fuscus) were modeled to create a "normal" echolocation signal for each bat which was then used as an artificial echo to synthesize a phantom target. The bat's task was to indicate which of two phantom targets (presented singly) was the "near" target and which the "far" target. Threshold range discrimination at a nominal target distance of 80 cm was about 0.6 cm for both bats. The normal signal was then modified to change the relative energy in each harmonic, the signal duration, the curvature of the frequency sweep, the absolute frequency, the phase of the second and third harmonics relative to the first, or the Doppler shift of the signal. To determine which modifications affected ranging performance, the altered models were used in tests of range discrimination that were interleaved on a day-to-day basis with tests using the normal model. Of the 12 modifications tested, only those changing the curvature of the frequency sweep affected performance. This result appears not to be predicted by current models of echo processing in FM bats. Eptesicus may be able to compensate for certain types of distortions of a returning echo, an ability possibly related to Doppler tolerance or to the characteristics of the natural variation in a bat's emissions.

Animals↗